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Transcript
CLASSIFICATION & EVOLUTION
Humans classify almost everything, including each other. This habit can be quite useful. For
example, when talking about a car someone might describe it as a 4-door sedan with a fuel injected
V-8 engine. A knowledgeable listener who has not seen the car will still have a good idea of what it
is like because of certain characteristics it shares with other familiar cars. Humans have been
classifying plants & animals for a lot longer than they have been classifying cars, but the principle is
much the same. In fact, one of the central problems in biology is the classification of organisms on
the basis of shared characteristics (i.e., simplesiomorphies vs. synapomorphies). As an example,
biologists classify all organisms with a backbone as "vertebrates." In this case the backbone is a
characteristic that defines the group. If, in addition to a backbone, an organism has gills & fins it is
a fish, a subcategory of the vertebrates. This fish can be further assigned to smaller & smaller
categories down to the level of the species. The classification of organisms in this way aids the
biologist by bringing order to what would otherwise be a bewildering diversity of species. (There
are probably several million species - of which about one million have been named and classified.)
The field devoted to the classification of organisms is called Taxonomy [Gk. taxis, arrange, put in
order + nomos, law].
The modern taxonomic system was devised by Carolus Linnaeus (1707-1778). It is a hierarchical
system since organisms are grouped into ever more inclusive categories from species up to
kingdom. Figure 1 illustrates how four species are classified using this taxonomic system. (Note
that it is standard practice to underline or italicize the genus & species names.)
KINGDOM
PHYLUM
CLASS
ORDER
FAMILY
GENUS
SPECIES
Animalia
Chordata
Mammalia
Primate
Carnivora
Hominidae
Canidae
Homo
Canis
sapiens
lupus
(human)
(wolf)
Arthropoda
Insecta
Hymenoptera
Apidae
Apis
mellifera
(honeybee)
Plantae
Angiospermophyta
Monocotyledoneae
Liliales
Liliaceae
Alium
sativum
(garlic)
Figure 1
In the 18th century most scientists believed that the Earth and all the organisms on it had been
created suddenly in their present form as recently as 4004 BC. According to this view, Linnaeus'
system of classification was simply a useful means of cataloging the diversity of life. Some
scientists went further, suggesting that taxonomy provided insight into the Creator's mind ("Natural
Theology").
This view of taxonomy changed dramatically when Charles Darwin published On The Origin of
Species in 1859. In his book Darwin presented convincing evidence that life had evolved through
the process of Natural Selection. The evidence gathered by Darwin, & thousands of
other biologist since then, indicates that all organisms are descended from a common ancestor. In
the almost unimaginable span of time since the first organisms arose (about 3.5 billion years) life
has gradually diversified into the myriad forms we see today.
As a consequence of Darwin's work it is now recognized that taxonomic classifications are actually
reflections of evolutionary history. For example, Linnaeus put humans & wolves in the class
Mammalia within the phylum Chordata because they share certain characteristics (e.g. backbone,
hair, homeothermy, etc.). We now know that this similarity is not a coincidence; both species
inherited these traits from the same Common Ancestor. In general, the greater the resemblance
between two species, the more recently they diverged from a common ancestor. Thus when we say
that the human & wolf are more closely related to each other than either is to the honeybee we mean
that they share a common ancestor that is not shared with the honeybee.
Wolf
Figure 2
The vertical axis in this figure represents time. The point at which two lines separate indicates
when a particular lineage split. For example, we see that mammals diverged from reptiles about 150
million years ago. The most recent common ancestor shared by mammals & reptiles is indicated
by the point labeled A. The horizontal axis represents, in a general way, the amount of divergence
that has occurred between different groups; the greater the distance, the more different their
appearance. Note that because they share a fairly recent ancestor, species within the same
taxonomic group (e.g. the class Mammalia) tend to be closer to each other at the top of the tree than
they are to members of other groups.
Several types of evidence can elucidate the evolutionary relationship between organisms, whether in
the form of a taxonomic classification (Fig. 1) or a tree (Fig. 2). One approach, as already
discussed, is to compare living species. The greater the differences between them, the longer ago
they presumably diverged. There are, however, pitfalls with this approach. For example, some
species resemble each other because they independently evolved similar structures in response to
similar environments or ways of life, not because they share a recent common ancestor. This is
called Convergent Evolution because distantly related species seem to converge in appearance
(become more similar). Examples of convergent evolution include the wings of bats, birds &
insects, or the streamlined shape of whales & fish. At first glance it might appear that whales are a
type of fish. Upon further examination it becomes apparent that this resemblance is superficial,
resulting from the fact that whales & fish have adapted to the same environment. The presence of
hair, the ability to lactate & homeothermy clearly demonstrate that whales are mammals. Thus, the
taxonomist must take into account a whole suite of characteristics, not just a single one.
The fossil record can also be helpful for constructing phylogenetic trees. For example, bears were
once thought to be a distinct group within the order Carnivora. Recently discovered fossils,
however, show that they actually diverged from the Canidae (wolves, etc.) fairly recently. The use
of fossils is not without its problems, however. The most notable of these is that the fossil record is
incomplete. This is more of a problem for some organisms than others. For example, organisms
with shells or bony skeletons are more likely to be preserved than those without hard body parts.
The Classification & Evolution of Artificial Organisms
In this exercise you will develop a taxonomic classification & phylogenetic tree for a group of
imaginary organisms. On the last page of this handout there are pictures of the 14 "living" & 58
"fossil" species that you will use. Take a look at the pictures & note the variety of appendages, shell
shapes, patterns, etc. Each species is identified by a number rather than a name. For fossil taxa
there is also a number in parentheses indicating the geological age of each specimen in millions of
years. Most of the fossils are extinct, but you will notice that a few are still living (e.g. species #24
is found among the living forms, but there is also a 2 million year old fossil of #24 in our
collection).
The purpose of this exercise is to illustrate the principles of classification & some of the processes
of evolution (e.g. Convergent Evolution). We do these exercises with artificial organisms so that
you will approach the task with no “preconceived” notion as to how they should be classified. This
means that you will have to deal with problems such as convergent evolution just as a taxonomist
would. With real organisms you would probably already have a pretty good idea of how they should
be classified & thus miss some of the benefit of the exercise.
Exercise 1: The Taxonomic Classification of Living Taxa
Carefully examine the fourteen living species & note the many similarities & differences between
them. Create a hierarchical classification of these species, using the format in Figure 3. Instead of
using letters (A, B, ...), as in this example, use the number of each species. Keep in mind that
Figure 3 is just a hypothetical example. Your classification may look quite different than this one.
GENUS 1
A
G
PHYLUM
CLASS 1
ORDER 1
FAMILY 1
FAMILY 2
GENUS 2
GENUS 3
GENUS 4
H
D
B
J
L
CLASS 2
ORDER 2
ORDER 3
FAMILY 3
FAMILY 3
GENUS 5
GENUS 6
E K C
F
I
Figure 3
The first step in this exercise is to decide which species belong in the same genus. Species within
the same genus share characteristics not found in any other genera. The taxa numbered 19 & 20 are
a good example; they are clearly more similar to each other than either is to any of the other living
species so we would put them together in their own genus. Use the same procedure to combine the
genera into families. Again, the different genera within a family should be more similar to each
other than they are to genera in other families. Families can then be combined into orders, orders
into classes & so on.
Exercise 2. The Comparative Approach to Phylogenetic Analysis
Construct a tree based only on your examination of the 14 living species. This tree
should reflect your taxonomic classification. For example, let us say you have put
species A & G into the same genus because you think they evolved from a common
ancestor (x). Their part of the tree would look like the diagram on the right.
A
When there are three or more species in a genus you must decide which two of the
E
species share a common ancestor not shared by the other(s). This diagram indicates
that species E & K are more closely related to each other than either is to C. We
y
hypothesize that E & K have a common ancestor (y) that is not shared by C. Similarly,
two genera that more closely resemble each other than they do other genera presumably
share a common ancestor. Thus, even in the absence of a fossil record it is possible to develop a
tree. We can even infer what a common ancestor like y might have looked like.
G
x
K C
z
Exercise 3. The Tree
Construct a tree for these taxa. Beginning with the oldest fossils, arrange the taxa according to their
evolutionary relationship. For your reconstruction, use the living taxa (remember some may be
contained in the fossil record) as well as taxa 15, 33, 36, 43, 44, 50, 52, 54, 57, 60, 73, 74, & 77.
*For extra credit, try reconstructing a tree using the entire data set.
Hints, Suggestions, & Warnings
a.
Draw lines to indicate the path of evolution. Start with the oldest fossil & work your way to the
present i.e., use a timescale as your “Y” axis.
b.
Branching should involve only two lines at a time:
Like this
YES
Not this
NO
c.
Some living forms are also found in the fossil record!
d.
There are gaps in the fossil record for some lineages. Also, some species went extinct without
leaving any descendants (this happens quite often in “real life”).
e.
The taxa were numbered at random; the numbers provide no clues to their evolutionary
relationships.
f.
There is only one correct phylogenetic tree in this exercise. Start with the most primitive form
(#73) & gradually modified it using a process that mimics evolution in real organisms.
Group Questions
1. Provide answers to Exercises 1 & 2.
2. Provide a diagram describing the relationship among your living & fossil Taxa.
3. Provide a justification as to why you constructed the tree the way you did i.e., are all your
groups, Genera & Families, Monophyletic? What characters were important in your
analysis?
4. What sorts of problems were encountered i.e., are there any regions of your tree that need
additional fossil data to reconstruct the relationships?
5. Did any of the taxa go extinct? How do they contribute to your understanding of the
Evolution of these species?
6. Are there any species, or groups, that show Convergent Evolution? Did these hinder your
efforts to contruct a tree, why or why not?
7. Would you describe the process of classification as difficult or easy, explain your answer?
How much more difficult would your job have been if you had been working from the
present to the past?
LIVING Taxa
FOSSIL Taxa
(numbers in parentheses indicate age in millions of years)